Yamna/Afanasevo elite males dominated by R1b-L23, Okunevo brings ancient Siberian/Asian population

afanasevo-okunevo

Open access paper New genetic evidence of affinities and discontinuities between bronze age Siberian populations, by Hollard et al., Am J Phys Anthropol. (2018) 00:1–11.

NOTE. This seems to be a peer-reviewed paper based on a more precise re-examination of the samples from Hollard’s PhD thesis, Peuplement du sud de la Sibérie et de l’Altaï à l’âge du Bronze : apport de la paléogénétique (2014).

Interesting excerpts:

Afanasevo and Yamna

The Afanasievo culture is the earliest known archaeological culture of southern Siberia, occupying the Minusinsk-Altai region during the Eneolithic era 3600/3300 BC to 2500 BC (Svyatko et al., 2009; Vadetskaya et al., 2014). Archeological data showed that the Afanasievo culture had strong affinities with the Yamnaya and pre-Yamnaya Eneolithic cultures in the West (Grushin et al., 2009). This suggests a Yamnaya migration into western Altai and into Afanasievo. Note that, in most current publications, “the Yamnaya culture” combines the so-called “classical Yamnaya culture” of the Early Bronze Age and archeological sites of the preceding Repin culture in the middle reaches of the Don and Volga rivers. In the present article we conventionally use the term Yamnaya in the same sense, in which case the beginning of the “Yamnaya culture” can be dated after the middle of the 4th millennium BC, when the Afanasievo culture appeared in the Altai.

Because of numerous traits attributed to early Indo-Europeans and cultural relations with Kurgan steppe cultures, members of the Afanasievo culture are believed to have been Indo-European speakers (Mallory and Mair, 2000). In a recent whole-genome sequencing study, Allentoft et al. (2015) concluded that Eastern Yamnaya individuals and Afanasievo individuals were genetically indistinguishable. Moreover, this study and one published concurrently by Haak et al. (2015) analyzed 11 Eastern Yamnaya males and showed that all of them belonged to the R1b1a1a (formerly R1b1a) (…)

indo-european-uralic-migrations-afanasevo
Early Chalcolithic migrations ca. 3300-2600 BC.

Published works indicate that R1b was a predominant haplogroup from the late Neolithic to the early Bronze Age, notably in the Bell Beaker and Yamnaya cultures (Allentoft et al., 2015; Haak et al., 2015; Lee et al., 2012; Mathieson et al., 2015). Nearly 100% of the Afanasievo men we typed belonged to the R1b1a1a subhaplogroup and, for at least three of them, more precisely to the L23 (xM412) subclade. (…)

(…) our results therefore support the hypothesis of a genetic link between Afanasievo and Yamnaya. This also suggests that R1b was indeed dominant in the early Bronze Age Siberian steppe, at least in individuals that were buried in kurgans (possibly an elite part of the population). The geographical and temporal distribution of subhaplogroup R1b1a1a supports the hypothesis of population expansion from West to East in the Eurasian steppe during this period. It should however be noted that the Yamnaya burials from which the samples for DNA analysis were obtained (Allentoft et al., 2015; Haak et al., 2015; Mathieson et al., 2015) were dated within the limits of the Afanasievo period. Ancestors of both East Yamnaya and Afanasievo populations must therefore be sought in the context of earlier Eneolithic cultures in Eastern Europe. Sufficient Y-chromosomal data from such Eneolithic populations is, unfortunately, not yet available.

mtdna-ydna-afanasevo-okunevo
Mitochondrial- (A) and Y- (B) haplogroup distribution in studied populations

Okunevo and paternal lineage shift in South Siberia

Results obtained in the current study, from more than a dozen Okunevo individuals belonging to the earliest stage of Okunevo culture, that is the Uibat period (2500–2200 BC) (Lazaretov, 1997), suggest a discontinuity in the genetic pool between Afanasievo and Okunevo cultures. Although Y-chromosomal data obtained for bearers of the Okunevo culture showed that one individual carried haplogroup R1b, most Okunevo Y-haplogroups are representative of an Asian component represented by paternal lineages Q and NO1.

Okunevo carrier of Y-haplogroup Q1b1a-L54, which also supports this hypothesis (L54 being a marker of the lineage from which M3, the main Ameridian lineage, arose). Okunevo people could therefore be a remnant paleo-Siberian population with possible Afanasievo input, as suggested by the presence of the R1b1a1a2a subhaplogroup in one individual.

indo-european-uralic-migrations-afanasevo-late
Late Chalcolithic migrations ca. 2600-2250 BC.

Replacement of Asian Indo-European elite lineages by R1a

Published genetic data from the late Bronze Age Andronovo culture from the Minusinsk Basin (Keyser et al., 2009), the Sintashta culture from Russia (Allentoft et al., 2015) and the Srubnaya culture from the region of Samara (Mathieson et al., 2015), show that males did not belong to Y-haplogroup R1b but mostly to R1a clades: there appears to have been a change in the dominant Y-chromosomal haplogroup between the early and the late Bronze Age in these regions. Moreover, as described in Allentoft et al. (2015), the Andronovo and Sintashta peoples were closely related to each other but clearly distinct from both Yamnaya and Afanasievo. Although these results do not imply that Y-haplogroup R1b was entirely absent in these later populations, they could correspond to a replacement of the elite between these two main periods and therefore a difference in the haplogroups of the men that were preferentially buried.

indo-european-uralic-migrations-okunevo-andronovo
Early Bronze Age migrations ca. 2250-1750 BC.

Afanasevo and the Tarim Basin

The discovery, in the Tarim Basin, of well-preserved mummies from the Bronze Age allows for the construction of two hypotheses regarding the peopling of the Xinjiang province at this period. The “steppe hypothesis,” argues for a link with nomadic steppe herders (Hemphill and Mallory, 2004), possibly represented in this case by Afanasievo populations and their descendants (Mallory and Mair, 2000). However, newly published cultural data from the burial grounds of Gumugou (Wang, 2014) and Xiaohe (Xinjiang, 2003, 2007) shows material culture and burial rites incompatible with the Afanasievo culture. The earliest 14C date for Tarim Basin burials would place them at the turn of the 2nd millenium BC (Wang, 2013), 500 years after the Afanasievo period.

Instead, early Gumugou and Xiaohe burial grounds were contemporary with the start of the Andronovo period. Likewise, the Bronze Age population of the Xinjiang at Gumugou/Qäwrighul is not phenotypically closest to Afanasievo but to the Andronovo (Fedorovo) group of northeastern Kazakhstan and western Altai (Kozintsev, 2009). Our investigations demonstrate that Y-chromosomal lineage composition is also compatible with the notion that the ancient Tarim population was genetically distinct from the Afanasievo population. The only Y-haplogroup found by Li et al. (2010) in the Bronze Age Tarim Basin population was Y-haplogroup R1a, which suggests a proximity of this population with Andronovo groups rather than Afanasievo groups.

I don’t think these finds are much of a surprise based on what we already know, or need much explanation…

I would add that, once again, we have more proof that the movement of Okunevo and related ancient Siberian migrants from Central or North Asia will not be able to explain the presence of Uralic languages spread over North-East Europe and Scandinavia already during the Bronze Age.

Also interesting is to read in more peer-reviewed papers the idea of Late Indo-European speakers clearly linked to the expansion of patrilineally-related elite males marked by haplogroup R1b-L23, most likely since Eneolithic Khvalynsk/Repin cultures.

Related:

Olalde et al. and Mathieson et al. (Nature 2018): R1b-L23 dominates Bell Beaker and Yamna, R1a-M417 resurges in East-Central Europe during the Bronze Age

The official papers Olalde et al. (Nature 2018) and Mathieson et al. (Nature 2018) have appeared. They are based on the 2017 preprints at BioRxiv The Beaker Phenomenon And The Genomic Transformation Of Northwest Europe and The Genomic History Of Southeastern Europe respectively, but with a sizeable number of new samples.

Papers are behind a paywall, but here are the authors’ shareable links to read the papers and supplementary materials: Olalde et al. (2018), Mathieson et al. (2018).

NOTE: The corresponding datasets have been added to the Reich Lab website. Remember you can use my drafts on DIY Human Ancestry analysis (viz. Plink/Eigensoft, PCA, or ADMIXTURE) to investigate the data further in your own computer.

olalde_pca
Image modified by me, from Olalde et al (2018). PCA of 999 Eurasian individuals. Marked is the late CWC outlier sample from Esperstedt, showing how early East Bell Beaker samples are the closest to Yamna samples.

I don’t have time to analyze the samples in detail right now, but in short they seem to convey the same information as before: in Olalde et al. (2018) the pattern of Y-DNA haplogroup and steppe ancestry distribution is overwhelming, with an all-R1b-L23 Bell Beaker people accompanying steppe ancestry into western Europe.

EDIT: In Mathieson et al. (2018), a sample classified as of Ukraine_Eneolithic from Dereivka ca. 2890-2696 BC is of R1b1a1a2a2-Z2103 subclade, so Western Yamna during the migrations also of R1b-L23 subclades, in contrast with the previous R1a lineages in Ukraine. In Olalde et al. (2018), it is clearly stated that of the four BB individuals with higher steppe ancestry, the two with higher coverage could be classified as of R1b-S116/P312 subclades.

This is compatible with the expansion of Indo-European-speaking Yamna migrants (also mainly of R1b-L23 subclades) into the East Bell Beaker group, as described with detail in Archaeology (and with the population movement we are seeing having been predicted) first by Volker Heyd in 2007.

yamna-bell-beaker
Yamna – East Bell Beaker migration 3000-2300 BC. Adapted from Harrison and Heyd (2007), Heyd (2007)

Also, the resurge of R1a-Z645 subclades in Czech and Polish lands (from previous Corded Ware migrants) accompanying other lineages indigenous to the region – seems to have happened only after the Bell Beaker expansion into these territories, during the Bronze Age, probably leading to the formation of the Balto-Slavic community, as I predicted based on previous papers. The fact that a sample of R1b-U106 subclade pops up in this territory is interesting from the point of view of a shared substrate with Germanic, as is the earlier BB sample of R1b-Z2103 for its connection with Graeco-Aryan dialects.

All this suggests that a North-West Indo-European dialect – ancestor of Italo-Celtic, Germanic, and Balto-Slavic -, supported in Linguistics by most modern Indo-European schools of thought, expanded roughly along the Danube, and later to northern, eastern, and western Europe with the Bell Beaker expansion, as supported in Anthropology by Mallory (in Celtic from the West 2, 2013), and by Prescott for the development of a Nordic or Pre-Germanic language in Scandinavia since 1995.

copper-age-late-bell-beaker
Diachronic map of Late Copper Age migrations including Classical Bell Beaker (east group) expansion from central Europe ca. 2600-2250 BC

Maybe more importantly, the fact that only Indo-Iranian-speaking Sintashta-Petrovka (and later Andronovo) cultures were clearly associated with R1a-Z645 subclades, and rather late – after mixing with early Chalcolithic North Caspian steppe groups (mainly East Yamna and Poltavka herders of R1b-L23 subclades) – gives support to the theory that Corded Ware (and probably the earlier Sredni Stog) groups did not speak or spread Indo-European languages with their migration, but most likely Uralic – as seen in recent papers on the much later arrival of haplogroup N1c – (compatible with the Corded Ware substrate hypothesis), adopting Indo-Iranian by way of cultural diffusion or founder effect events.

As Sheldon Cooper would say,

Under normal circumstances I’d say I told you so. But, as I have told you so with such vehemence and frequency already the phrase has lost all meaning. Therefore, I will be replacing it with the phrase, I informed you thusly

I informed you thusly: